Design, function and evolution in the wings of holometabolous insects

The wings of Holometabola are reviewed in functional terms. Two basic types, anteroposteriorly symmetric (‘symmetric’) and anteroposteriorly asymmetric (‘asymmetric’), are recognized and their modes of operation discussed. Most neuropteroids have asymmetric wings, whereas relative anteroposterior symmetry appears plesiomorphic for mecopteroids. Wing coupling has only occurred in lineages with wings of symmetric type, but anteroposterior asymmetry of the wing couple has later developed several times in association with improved, more versatile, flight performance. Coleoptera wings seem to have developed from predecessors of symmetric type, but their morphology is strongly influenced by the need to fold up at rest. Diptera provide an excellent illustration of the evolution of asymmetric, high‐performance wings from the symmetric, ‘mecopteran’ type. Asymmetric and symmetric wings can both be derived from less specialized types. Both are well represented as fossils from the Permian onwards, and these supply further examples of the evolution of asymmetry from the symmetric pattern.